Official journal website: Amphibian & Reptile Conservation amphibian-reptile-conservation.org 9(2) [Special Section]: 1–14 (e106).

The anuran fauna of a West African urban area

1,*N’Goran Germain Kouamé, 2,3Caleb Ofori-Boateng, 2,3Gilbert Baase Adum, 4Germain Gourène, and 5,†Mark-Oliver Rödel

1Jean Lorougnon Guédé University, Department of Biology and Animal Physiology, UFR-Environnement, Daloa, BP 150 CÔTE D’IVOIRE 2Forestry Research Institute of Ghana, P.O. Box 63, Fumesua, Kumasi GHANA 3Department of Wildlife and Range Management, Faculty of Renewable Natural Resources, Kwame Nkrumah University of Science and Technology, Kumasi GHANA 4Nangui Abrogoua University, Laboratoire d’Environnement et de Biologie Aquatique, UFR-SGE, 02 BP 801, Abidjan 02 CÔTE D’IVOIRE 5Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstrasse 43, 10115 Berlin GERMANY

Abstract.—Reported are the results of an amphibian survey in the district of Daloa and surroundings, in central-western Ivory Coast. Spanning a three year period, we investigated two general areas, each during the rainy and dry seasons. During 62 days of field work 30 anuran species were recorded. The urban environment mainly contained widespread anuran species with preferences for savannah-dominated landscapes and farmbush habitats. The recorded total anuran species richness in the urban area exceeded the diversity in the savannah islands/forest mosaic bordering the Daloa district. This indicates many savannah species may do well in urban situations. However, this higher species richness was only due to one site that possessed particularly diverse amphibian breeding sites, thus illustrating the necessity of maintaining suitable habitats for a wide-range of species. One of the most surprising findings was Kassina schioetzi, a species usually difficult to find in its natural habitat. In Daloa it seems to have successfully adapted to the urban conditions. Although the anuran richness in the Daloa area was relatively low compared to other Ivorian humid savannah areas, it supported an important part of the countries amphibian diversity. Nevertheless the forest habitats, and specifically the forest islands bordering the Daloa district, should be considered sensitive conservation areas.

Key words. Amphibians, conservation status, Côte d’Ivoire, Upper Guinea, urban ecology

Citation: Kouamé NG, Ofori-Boateng C, Adum GB, Gourène G, Rödel MO. 2015. The anuran fauna of a West African urban area. Amphibian & Reptile Conservation 9(2) [Special Section]: 1–14 (e106).

Copyright: © 2015 Kouamé et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial- NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website .

Received: 16 June 2015; Accepted: 29 October 2015; Published: 16 November 2015 West Africa has been experiencing intensive urbaniza- demand for agriculture and urbanization (Bible 2013; tion to such a point that human modified landscapes are Hansen et al. 2013). gradually taking over the majority of natural landscapes, The Haut-Sassandra region is traditionally an im- in particular native forests (Deikumah and Kudom 2010; portant trading center, particularly for cocoa production Bible 2013). Whereas various Ghanaian forests are pro- in the Ivory Coast, has attracted 44.8% of national and tected and/or sustainably managed (Adum et al. 2013; 23.4% of foreign farmers (Assiri et al. 2009). During the Ofori-Boateng et al. 2013), only very few Ivorian for- country’s 2010‒2011 post election violence, Daloa, the est remnants receive sufficient protection and sustainable third largest city of the country and the regional capital management (e.g., see Mayaux et al. 2004). The Ivorian of the Haut-Sassandra region, became a refuge for peo- population has exploded over the past four decades, tri- ple from the northern, central, and western Ivory Coast, pling from 6.7 million in the early 1970s to approxi- resulting in a rapid urbanization process. As one result, mately 22 million people today (Bible 2013), has largely forests surrounding the city are increasingly fragmented. accelerated an urbanization process causing massive en- To enhance the protection of biological diversity, the vironmental damage. The gradual disruption of forests, Ivorian Ministry of Scientific Research has therefore re- has worsened in several forested areas of the Ivory Coast cently recommended the collection of scientific informa- during the prolonged political crisis in the first decade of tion to update the biodiversity data of the Haut-Sassandra the 21st century, has mainly stemmed from increase land region. As data for amphibians were still lacking, we sur- Correspondence. Emails: *[email protected]; †[email protected] (corresponding authors).

Amphib. Reptile Conserv. 1 November 2015 | Volume 9 | Number 2 | e106 Kouamé et al. veyed the amphibian fauna within the district of Daloa other infrastructure. We surveyed two general areas: 1) and its surroundings, and herein report for the first time the district of Daloa (urban area), and 2) the savannah an assessment of the species richness and composition of islands/forest mosaic bordering the Daloa district (non- the anuran fauna in a West African urban area. urban aspect). Our surveys were covering a three year period (see Appendix 1 for further details). We inves- Methods tigated four sites inside the urban area namely: Balou- zon (Bal: 45 ha), Eveché (Eve: 50 ha), Gbokora (Gbo: Study area. Daloa is the third largest city in the 80 ha), and Tazibouo (Taz: 100 ha). As a comparison, Ivory Coast and the regional capital of the Haut- we surveyed Sapia (Sap: 150 ha) and Zaibo (Zai: 190 Sassandra region. It is situated in central-west- ha), two non-urban sites in the savannah islands/forest ern Ivory Coast (06°53’01.8”‒06°94’97.8” N; mosaic adjacent to urban Daloa (see Appendix 1). The 006°25’65.3”‒006°68′89.0” W), in the transition zone Balouzon and Eveché areas were mainly characterized between semi-deciduous forest and humid Guinea savan- by unpaved roads, a swampy area used for vegetable cul- nah. The town is an important trading center, particularly tivation, and a concentration of buildings. A large stream, for cocoa. The region has a mean annual temperature bordered by coconut trees and grasses, was used for fish- of 26.3 °C; the annual precipitation ranges from 1,200 ing activities. The vegetation in Gbokora was dominated to 1,700 mm. The climate includes a long rainy season by grasses and a semi-deciduous forest interspersed by (April to June) with the highest precipitation peak in a highway. Some swampy areas surveyed in this site June, a short dry season (July to August) alternating with were being used for vegetable cultivation. This area was a short rainy season (September to October), and a long noisy due to heavy traffic. A concentration of buildings dry season (November to March). The relative mean hu- and streetlights were characteristic of the Gbokora site. midity is 75% (Eldin 1971). The Tazibouo site mainly consisted of unpaved roads, Description of the survey areas (Fig. 1). Our defini- buildings, semi-deciduous forest adjacent the Daloa Jean tion of an urban area follows McDonnell and Pickett Lorougnon Guédé University, and some construction (1993) and Demographia (2008), i.e., taking into con- sites. This site also comprised the Theological and Pas- sideration a minimum density of 400 humans/km2 and toral Institute of Daloa whose garden was dominated by other factors such as density of buildings, roads and bamboo, other grasses, and several stands of different or-

Fig. 1. Typical aspects of habitats of the urban Daloa; a = concentration of houses in a high population zone; b = degraded forest on the periphery of the urban area; c = highway crossing degraded forest and farmbush; d = amphibian breeding pond in the urban environment.

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namental plants. Some water bodies, i.e., two large per- ogy or threats, or being particularly typical for the urban manent ponds situated near a roadside and bordered by amphibian fauna. The nomenclature used herein follows grasses, were encountered. The ponds served as a water the taxonomy by Frost (2015). After capture, frogs were point for cattle. A few termite mounds were also present. identified to species level, measured, sexed, and ifnot A swampy area investigated was used for rice and veg- kept as vouchers, released in their respective habitats. etable cultivations. Some parts of this area were light- Snout-urostyle-length (SUL) was taken with a dial cali- ened by streetlights at night. The sites Sapia and Zaibo, per (accuracy ± 0.5 mm). Records of Xenopus muelleri adjacent to the district of Daloa represented non-urban were based on visual observations only. For all other spe- conditions. However, they had lost the majority of their cies we deposited vouchers at the Jean Lorougnon Guédé natural forest cover, resulting in an overall change from University, Daloa, Ivory Coast (see Appendix 3). Frog a forest to a savannah-dominated landscape. Both sites vouchers were euthanized humanly in a 1,1,1-Trichloro- mainly consisted of farmbush, small farms, coffee, and 2-methyl-2-propanol hemihydrate (MS222) solution and cocoa plantations. Some swamps that were part of these thereafter preserved in 70% ethanol. two sites had been converted to rice fields. Small forest Statistics. We used the daily species lists to calculate islands were still encountered at Zaibo, but fewer forest the sampling efficiency. We calculated the estimated spe- islands were left at the Sapia site. cies richness with the Chao2 and Jack-knife1 estimators Field work, sampling effort and vouchers. Amphib- (software: EstimateS, Colwell 2006). These estimators ians were mainly located opportunistically, during visual are incidence based, calculating with the presence/ab- and acoustic surveys of all available habitats by NGK. sence data of the daily species lists (62 days of survey Surveys were undertaken daily between 07:00‒11:00 work) for 30 anuran species. To avoid order effects we and 18:00‒22:00 GMT over a total of 62 days (see Ap- accomplished 500 random runs of the daily species lists. pendix 2) at all general survey areas. A hand-held GPS The Sørensen’s Similarity Index (β) was used to deter- receiver (Garmin 12XL) was used to record the geo- mine the extent of similarity between the two main sur- graphical positions of all study sites. The searching tech- veyed areas (herein the district of Daloa and the savan- niques used included acoustic surveying, visual scanning nah islands/forest mosaic bordering the Daloa district; β of terrain and refuge examination (e.g., lifting logs and may vary from 0 to 1 (Sørensen 1948; Wolda 1981). rocks, peeling away barks, scraping through leaf litter, looking around or within burrows, and termite mounds). Results Amphibians encountered were not marked and repeated sightings thus cannot be excluded. As we only include Species richness and faunal similarities presence/absence data and not abundances in our analy- ses this seem to be of negligible importance. In total we recorded 30 anuran species (Table 1). Acous- Below we comment only on a few species being re- tics indicated more than one Arthroleptis species live in markable concerning their distribution, taxonomy, biol- our area. So far, it is not possible to separate taxa from

Table 1. Anuran species recorded in the urban and non-urban areas of Daloa, with sites (see Appendices 1–3), general habitat prefer- ence and distribution range. S = savannah, FB = farmbush (degraded forest and farmland), F = forest, A = Africa (occur also outside West Africa), WA = West Africa (defined as the area west of the Cross River in Nigeria), UG = Upper Guinea (forest zone west of the Dahomey Gap), E = endemic to Ivory Coast and eastern Guinea, * = taxon comprise complex of several species, ** = records on this survey comprise several species (according to acoustics). Family / Species Site Habitat Distribution S FB F A WA UG E Arthroleptidae Arthroleptis spp.** Bal, Eve, Gbo, Sap, Taz, Zai — X X — — X (?) Leptopelis spiritusnoctis Zai X X — X — — L. viridis Bal, Eve, Gbo, Taz X — — X — — — Bufonidae Amietophrynus maculatus Gbo, Sap, Taz, Zai X X — X — — — A. regularis Taz X X — X — — — Dicroglossidae Hoplobatrachus occipitalis Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — — Hemisotidae Hemisus marmoratus Taz, Zai X X — X — — — Hyperoliidae Afrixalus dorsalis Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — —

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Table 1 (continued). Anuran species recorded in the urban and non-urban areas of Daloa, with sites (see Appendices 1–3), general habitat preference and distribution range. S = savannah, FB = farmbush (degraded forest and farmland), F = forest, A = Africa (occur also outside West Africa), WA = West Africa (defined as the area west of the Cross River in Nigeria), UG = Upper Guinea (forest zone west of the Dahomey Gap), E = endemic to Ivory Coast and eastern Guinea, * = taxon comprise complex of several species, ** = records on this survey comprise several species (according to acoustics).

Family / Species Site Habitat Distribution Hyperoliidae (cont.) S FB F A WA UG E Hyperolius concolor concolor Bal, Eve, Gbo, Sap, Taz, Zai X X — — — X — H. fusciventris fusciventris Sap X X — — X — H. guttulatus Zai X X — X — — — H. nitidulus Bal, Eve, Gbo, Sap, Taz X — — — X — — H. picturatus Gbo, Sap, Taz, Zai — X X — — X — H. sp. Taz — X X — — X — Kassina schioetzi Taz X X — — — X — K. senegalensis Sap, Taz X — — X — — — Phrynobatrachidae Phrynobatrachus calcaratus* Sap — X X X — — — P. francisci Bal, Taz X — — — X — — P. gutturosus* Sap, Taz, Zai X X X — — X — P. latifrons Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — — Phrynomeridae Phrynomantis microps Taz X — — X — — — Pipidae Xenopus muelleri Taz X — — X — — — Ptychadenidae Ptychadena bibroni Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — — Ptychadena mascareniensis* Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — — Ptychadena oxyrhynchus Sap, Taz, Zai X X — X — — — Ptychadena tournieri Sap, Taz X — — — X — — Ptychadena pumilio Bal, Eve, Gbo, Sap, Taz, Zai X X — X — — — Ptychadena tellinii Taz X — — X — — — Ranidae Amnirana albolabris Sap, Zai — X X X — — — A. galamensis Taz X — — X — — —

the Arthroleptis poecilonotus-complex based on mor- richness. More than one fifth of the encountered species phology. They can be distinguished by advertisement call (seven spp., 23%; Table 1) depend on forest but tolerate and genetic characters. However, assigning populations, farmbush habitats (degraded forest). Nine species (30%) based on these characters, to available names (indistin- are very closely associated with savannah habitats. Thir- guishable museum types without molecular data) is not teen species (43%) exhibit a strong preference for savan- possible (for a short review of the taxonomic situation in nah and farmbush habitats and are normally not found West African Arthroleptis spp. see Rödel and Bangoura in forest. Four species (13%) do not occur outside West 2004). We thus provisionally lumped all records of this Africa [defined as the area west of the Cross River in Ni- genus as Arthroleptis spp. A list of recorded anurans with geria; see Penner et al. (2011)], and are often restricted to site records, known habitat preference and their distribu- smaller parts of West Africa. Seven of all recorded spe- tion ranges is given in Table 1. cies (23%) occur only in the Upper Guinea forest zone Based on the daily species lists we calculated our (forests west of the Dahomey Gap). The total number of sampling efficiency. The Jack-knife 1 estimator - calcu species recorded in the district of Daloa was 25, while the lated 33 anuran species, the Chao 2 estimator estimated species richness in the adjacent savannah/forest mosaic 31 species for the study area. We hence recorded almost was 21. However, the high species number for Daloa was the entire (94% and 99%, respectively) estimated species mainly due to one site (Tazibouo). When excluding this

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Table 2. Sørensen’s similarity values for pairwise comparisons highest abundances directly around human settlements. of the anuran community between the six surveyed sites (see At night, it was found in gardens, around houses, park- text and Appendix 1). ing areas, buildings, or below streetlights, preying mostly Sites Eve Gbo Taz Sap Zai on insects. During the day, it was found under rocks or Bal 0.95 0.87 0.61 0.62 0.59 logs. The most imminent threat to the toad’s survival in the city of Daloa is its exploitation for scientific courses Eve – 0.91 0.57 0.64 0.61 at the university. Every year several hundred individu- Gbo – – 0.65 0.67 0.71 als are collected by students and subsequently killed and Taz – – – 0.65 0.63 dissected in anatomy courses. This exploitation seems to Sap – – – – 0.71 have reached a point where the species is becoming rare in the city. However, concerning the entire range of the site diversity was higher in the savannah/forest mosaic. species, it is very common and of Least Concern (IUCN The number of species common to both areas was 16 2015). (Sørensen’s Similarity Index β: 0.70). Within the district Hoplobatrachus occipitalis (Günther, 1859) is the of Daloa we recorded 11 species in Balouzon, 10 species most commonly consumed frog species in West Africa. in Eveché, 12 in Gbokora, and 25 in Tazibouo. Within the The frog trade varies regionally from e.g., local scale in savannah/forest mosaic we recorded 18 and 16 species in Burkina Faso, to intensive cross-border trade in north- Sapia and Zaibo, respectively. ern Benin and Nigeria (Mohneke et al. 2009, 2010). The The results of the Sørensen’s similarity for pairwise consumption of H. occipitalis (Fig. 3a) has recently in- comparisons in the six surveyed sites are presented in creased to a considerable extent in the Ivory Coast where Table 2. At least more than 50% of the recorded species this species is an important component of animal protein were similar between sites. The anuran fauna of Daloa in some local populations (NGK, unpubl. obs.). In Daloa, urban area was most similar to that of the Comoé Na- the trade of H. occipitalis mainly took place on a local tional Park, a savannah area in northern Ivory Coast (β: scale at the different markets of the district. Usually a 0.72). With 68% and 66% faunal similarity the Lamto batch of five adult specimens was sold for 500.00 FCFA Faunal Reserve and the Marahoué National Park, which (app. 0.84 USD). Frog meat are sold fresh (Figs. 3b, c) are situated in the same vegetation zone as Daloa, were or dried (Fig. 3d). It is used in soups, stews, or sauces by very similar to the Daloa fauna. Other Ivorian protected the local populations. The local price in Daloa markets areas such as the Mont Péko and Mont Sangbé National was mean to low compared to prices recorded in Burkina Parks comprise savannah and real rainforest zones and Faso and Nigeria, respectively. According to Mohneke thus consequently differed in their faunal composition, et al. (2010), in Burkina Faso, the price for one frog de- compared to Daloa (Table 3). pended on its size and varied between 25.00 FCFA for a small frog, up to 250.00 FCFA (0.05 USD and 0.50 Species accounts USD) for a large one. In Nigeria, they reported one bag containing at least 1,000 frogs cost 26.94–40.40 USD on Amietophrynus regularis (Reuss, 1833) ‒ The genus purchase and 40.40–67.34 USD at sale. In the urban area Amietophrynus currently encompasses 40 species of true of Daloa hard data on harvested frog numbers and re- African toads [Frost 2015; although this list also con- spective consequences for the local populations are lack- tains non-vaild taxa such as Amietophrynus chudeaui ing. The local trade of H. occipitalis hence needs more (Chabanaud, 1919) see Rödel (2000)]. Amietophrynus attention and detailed investigation. regularis has a wide distribution in Africa and inhabits Hyperolius concolor concolor (Hallowell, 1844)(Fig. a broad range of habitats from moist and dry savannahs, 4) is a typical West African farmbush species living in montane grassland, forest margins, and agricultural habi- degraded forest of the forest zone and gallery forests in tats, as well as human settlements, often in association the savannah zone (Schiøtz 1967; Rödel 2000). It seemed with rivers (Rödel 2000; Channing and Howell 2006). In to do very well under urban condition and was hence our urban sites A. regularis (Fig. 2) seemed to reach its among the most widespread species recorded in the ur-

Table 3. Sørensen’s similarity value (β) between the anuran fauna of the Daloa urban area and other Ivorian areas, and respective species richness; twenty-five species were recorded in urban Daloa (this study); NP= national park; FR= faunal reserve. Number of species common Area Species richness β-value (Sørensen) Source with the Daloa urban area Comoé NP 33 21 0.72 Rödel and Spieler (2000) Lamto FR 40 22 0.68 Adeba et al. (2010) Marahoué NP 33 19 0.66 Rödel and Ernst (2003) Mont Péko NP 33 11 0.38 Rödel and Ernst (2003) Mont Sangbé NP 45 20 0.57 Rödel (2003)

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Fig. 2. Amietophrynus regularis female recorded in the garden of the Theological and Pasto- ral Institute of Daloa. ban sites of Daloa (Table 1); it was particularly abundant (Rödel et al. 2002; Rödel and Ernst 2003; Adeba et al. among grasses near ponds. In the rainy season we record- 2010). It lives along the savannah forest edge, reaching ed some, presumably migrating, individuals on windows, into the savannah zone along rivers. The species may balconies, and in houses. also occur in Bia National Park, western Ghana, but a Hyperolius sp. ‒ The genus Hyperolius Rapp, 1842 is voucher from there exhibited a mixture of characters one of the most diverse African anuran genera with cur- with K. cochranae (Hillers et al. 2009). Kassina schioetzi rently approximately 28 species occurring in West Africa is usually hard to find in all localities so far investigated (Schiøtz 1967, 1999; Frost 2015). A major taxonomic (see above and own experience of the authors). In the problem is many species of this genus are highly variable district of Daloa (Tazibouo), some males were observed (e.g., Schiøtz 1999). On 15 September 2013, at around calling at night from more exposed sites (Fig. 6). We also 07h00 GMT, we found a Hyperolius on humid ground in encountered a small number of other males calling in a the Tazibouo site, within the garden of the Theological bamboo patch within the Theological and Pastoral Insti- and Pastoral Institute, after it had rained heavily the night tute, and at a grassy roadside in the vicinity of a large before. Our individual lacked a vocal sac and gland and pond. Our recorded males measured 32.1 ± 1.6 (SUL, n hence is either female or juvenile (Fig. 5). It resembles = 4), thus being within the known range of K. schioetzi either a juvenile H. picturatus or a newly metamorphosed (Rödel et al. 2002). individual phase J of H. sylvaticus ivorensis, which is Leptopelis viridis (Günther, 1868) (Fig. 7) is one of normally brownish to green with paired undelimited dor- the most characteristic species inhabiting the West Af- solateral stripes, and an hourglass pattern (Schiøtz 1999). rican savannahs and the degraded areas of the former The size of our reed frog was 20 SUL, thus exceeding the rainforest belt. As a synanthropic species, it also lives in size of freshly metamorphosed Hyperolius of most spe- villages (Schiøtz 1967; Rödel 2000). It is one of the most cies (compare e.g., Schiøtz 1967; Rödel 2000). Its dorsal widespread anurans in the urban sites of Daloa. Leptope- surface was beige with a greenish grey hourglass pattern. lis viridis was found around houses, and in gardens. The The iris was golden, the anterior and posterior sides of majority of the recorded males were found at night call- pupil were red. The ventral surface was whitish. Without ing exposed on the ground between short grasses, which having male specimens and advertisement calls available is in contrast to the calling sites in natural habitats. There it cannot be decided if this frog represents an undescribed the species calls, often from high perch sites, in bushes species or only an atypical, but known Hyperolius spe- and trees (Grafe et al. 2000; Rödel 2000). cies. Phrynomantis microps Peters, 1875 is a medium- Kassina schioetzi Rödel, Grafe, Rudolf, and Ernst, sized microhylid frog inhabiting the savannah regions of 2002 was known so far from the Mont Péko National West Africa (Hirschfeld and Rödel 2011) where it hides Park, the Marahoué and Comoé National Parks, and the in burrows or empty termite mounds during the day and Lamto Faunal Reserve, all situated in the Ivory Coast the dry season. The frog was also observed to occupy and

Amphib. Reptile Conserv. 6 November 2015 | Volume 9 | Number 2 | e106 The anuran fauna of a West African urban area live essentially unharmed in the nest of the highly ag- most 85% of amphibian species threatened by urbaniza- gressive ant species – Paltothyreus tarsatus (e.g., Rödel tion are encountered in the tropics (IUCN, Conservation and Braun 1999; Rödel et al. 2013). In Daloa, P. microps International and NatureServe 2006). Many factors are was heard calling at night in tufts of grass around houses known to negatively influence the herpetofauna inhabit- after heavy rainfalls. In the garden of the Theological and ing big cities. Among these factors are habitat loss, habi- Pastoral Institute, a calling male was observed in associa- tat fragmentation, isolation, pollution, over harvesting, tion with an Emperor Scorpion (Pandinus imperator) in and road traffic (Hammer and McDonnell 2008; Perry et a hole behind the wall of a building. The association of P. al. 2008; Stuart et al. 2008; Deikumah and Kudom 2010; microps with scorpions has also been reported by Rödel Tonini et al. 2011). However, many species are able to and Braun (1999) and Rödel (2000). We captured another adapt to urban conditions and sometimes urban areas male (Fig. 8) on 08 September 2013 around 22h00 GMT may even surprise with the discovery of scientifically at the edges of a wide roadside pond beside the Theologi- new species (Newman et al. 2012; Feinberg et al. 2014; cal and Pastoral Institute. Howlader et al. 2015). This also concerns the Ivorian Xenopus muelleri (Peters, 1844) is an aquatic species city of Abidjan where a monotypic genus Morerella cy- inhabiting the West African savannah ponds of highly anophthalma and a night-frog Astylosternus laticephalus variable size during the rainy season and the edges of riv- have recently been discovered and described (Rödel et ers during the dry season (Rödel 2000). In the urban site al. 2009, 2012). Tazibouo, the frog was observed to live in holes drilled With its geographic position in a transition zone be- in the ground by the national company of water distribu- tween the semi-deciduous forest and humid savannah, tion. The depths of these holes varied from 0.7‒1.20 m. we expected the urban landscape of Daloa region to pro- mote a diverse amphibian fauna. However, the overall Discussion species richness (30 spp.) was lower compared to the species richness recorded in western, central, and north- Despite their importance to ecosystem functions ern Ivorian savannah areas, for instance the Mont Sangbé (Mohneke and Rödel 2009; Hocking and Babbitt 2014), National Park (45 species, Rödel 2003), Lamto Faunal amphibians are still among the least studied vertebrates Reserve (40 species, Adeba et al. 2010), Marahoué and particularly in urban and suburban areas in the tropics Mont Péko National Parks (33 species for each park, (Hamer and McDonnell 2008; Pickett et al. 2001). Al- Rödel and Ernst 2003), or the Comoé National Park (33

Fig. 3. Hoplobatrachus occipitalis from the district of Daloa (a) and a woman trading this species on a local market (b); batches of five adult specimens, fresh (b and c) or dried (d), were sold for 500.00 FCFA (app. 0.84 USD).

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Fig. 4. A calling Hyperolius concolor concolor male recorded at the garden of the Theological and Pastoral Institute of Daloa.

Fig. 5. Dorsolateral view of a juvenile Hyperolius sp. with uncertain taxonomic status from the urban Daloa. species, Rödel and Spieler 2000). Compared to these and turosus (Rödel and Branch 2002; Assemian et al. 2006; other West African savannah areas with known amphib- Kouamé et al. 2014; Kpan et al. 2014; the latter two spe- ian assemblages such as north-western Benin (Nago et al. cies comprising out of cryptic species with savannah and 2006), east-central Guinea (Greenbaum and Carr 2005), forest specialists), most of the recorded frogs were wide- central-northern Guinea (Hillers et al. 2008a), or eastern spread species with preferences for savannah-dominated Ghana (Leaché et al. 2006), the urban landscape of Daloa landscape and farmbush habitats. The six surveyed sites ranks among the West African areas of medium to low all shared at least half of their species with all other sites. amphibian species richness. While we recorded few for- We observed the highest species richness at the Tazibouo est related species e.g., Amnirana albolabris, Leptopelis site (25 spp.) which was the only urban site comprising spiritusnoctis, Phrynobatrachus calcaratus, and P. gut- various suitable breeding habitats. For instance in the

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Fig. 6. Kassina schioetzi from Daloa urban area; shown is a male calling from the ground (a), and its whitish venter and almost circular gular gland (b) characteristic for the species. garden of the Theological and Pastoral Institute, ephem- eas, in the form of the collection of anurans for anatomy eral and perennial wetlands, well-suited for the co-ex- and food consumption. Compared to European towns istence of species with different reproductive strategies (e.g., Mollov 2005), however, there are still plenty of were present: e.g., very small temporary breeding sites habitats available to amphibians, the traffic is usually less (Phrynobatrachus spp.), larger, almost permanent breed- intense as many of the roads remain unpaved allowing ing sites (Afrixalus dorsalis, Hyperolius concolor, Kas- frogs to cross. In fact our non-urban sites were all within sina schioetzi, K. senegalensis, Phrynomantis microps), a matrix of agricultural land and thus most likely prone and humid places for species with terrestrial direct de- to a variety of pesticides which could be a higher threat velopment (Arthroleptis poecilonotus-group). This gar- than the threats experienced in towns. The adaptability of den also played an important role in providing refuge for amphibians within the urban development seemed to be several other species in particular during the heat of the species-specific and was highly variable even between day and the dry season. sites. For example some species such as Hyperolius In addition to the fact that many amphibian species guttulatus, H. fusciventris fusciventris, Amnirana albo- depend on different but complementary habitats (e.g., labris, Leptopelis spiritusnoctis, and Phrynobatrachus aquatic sites for the tadpoles, terrestrial site of the meta- calcaratus, encountered in the savannah-forest mosaic morphosed individuals), their populations are usually outside of Daloa were never found in the urban sites. This structured as meta-populations (Pope et al. 2000; Marsh is most likely due to the fact that their specific habitats and Trenham 2001). Urbanization and in particular frag- are no longer present. For instance Hyperolius guttulatus mentation and isolation of habitats by roads and other breeds almost exclusively in very large and deeper ponds urban infrastructure is reducing the connectivity of popu- (Rödel 2000; Schiøtz 1967, 1999); and Phrynobatrachus lation networks (Vos and Chardon 1998). Hence, we ex- calcaratus typically lives at rain forest edges or in gallery pected to record lower amphibian diversity in the district forests in the savannah zone (Rödel 2000). Respective of Daloa than in the savannah-forest mosaic adjacent to habitat types for both latter species were not found in the this district. Surprisingly, the total anuran richness in urban environment. It is known that in forested areas the the urban environment was higher than in the adjacent alteration of the microclimate, due to degradation of the savannah-forest mosaic. This result indicates many am- vegetation structure, causes a shift in species composi- phibian species may survive under urban situations, such tion (Ernst and Rödel 2005, 2006; Hillers et al. 2008b; as in the district of Daloa. However, this high total spe- Ofori-Boateng et al. 2013). Such effects might be even cies richness was due to only one of four urban sites, i.e., worse in the usually more open habitats of urban areas. the Theological and Pastoral Institute, comprising many different habitat types and particularly diverse breeding Conclusion sites. The other urban sites actually had slightly lower species richness than the non-urban sites. This illustrates The study is indicating that an unexpected high number a high amphibian diversity in urban areas may be main- of anuran species seem to be able to survive in a current tained and even exceed such as of nearby non-urban African city. However, this is not the case for all species. areas; however, this can only be achieved by offering a For those species the protection of natural forest and sa- wide range of different habitats suitable for various am- vannah ecosystems is very important. The forest habi- phibian species. tats, and specifically the forest ‘‘islands” bordering the Apart from roads potentially reducing or ceasing gene Daloa district, should thus be considered sensitive areas flow, amphibians further face direct threats in urban ar- and dispersal corridors need to be maintained. Within the

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Fig. 7. Dorsolateral view of Leptopelis viridis, one of the most Fig. 8. Dorsolateral view of a male Phrynomantis microps from widespread anurans from the Daloa urban area. Daloa urban area. urban areas, the availability of a diverse set of habitats species richness and shared species from samples. is a prerequisite for the maintenance of high amphibian Version 7.5.2. Available: http://Viceroy.eeb.uconn. species richness. edu/estimates (Accessed: 14 February 2010). Deikumah JP, Kudom AA. 2010. Biodiversity status of urban remnant forests in Cape Coast, Ghana. Journal Acknowledgments.—We are indebted to Dago Gna- of Science and Technology 30: 1‒8. kri, President of the Jean Lorougnon Guédé University, Demographia. 2008. World urban area and population for providing authorization to undertake this survey. We projects. Wendell Cox Consultancy, Belleville, Illi- thank Daplex H. Ouenchist, Director of the Theological nois. Available: www.demographia.com/db-worldua. and Pastoral Institute for permitting us to investigate the pdf (Accessed 08 July 2014). garden of his institution. We are particularly grateful for Eldin M. 1971. Le climat. Pp. 73–108 in: Le milieu na- the support and collaboration from Chief Nanan Kra, el- turel de la Côte d’Ivoire. Editors, Avenard JM, Eldin der of the Baoulé-Ayétou from the Haut-Sassandra re- M, Girard G, Sircoulon J, Touchebeuf P, Guillau- gion. met E, Adjanohoun E, Perraud A. ORSTOM, Paris, France. 401 p. Literature Cited Ernst R, Rödel MO. 2005. Anthropogenically induced changes of predictability in tropical anuran assem- Adeba PJ, Kouassi P, Rödel MO. 2010. Anuran amphib- blages. Ecology 86: 3,111–3,118. ians in a rapidly changing environment – revisiting Ernst R, Rödel MO. 2006. Community assembly and Lamto, Côte d’Ivoire, 40 years after the first herpeto- structure of tropical leaf-litter anurans. Ecotropica 12: faunal investigations. African Journal of Herpetology 113–129. 59: 1–18. Feinberg JA, Newman CE, Watkins-Colwell GJ, Adum GB, Eichhorn MP, Oduro W, Ofori-Boateng C, Schlesinger MD, Zarate B, Curry BR, Shaffer HB, Rödel MO. 2013. Two-stage recovery of amphibian Burger J. 2014. Cryptic diversity in Metropolis: Con- assemblages following selective logging of tropical firmation of a new leopard frog species (Anura: Ra- forests. Conservation Biology 27: 354–363. nidae) from New York City and surrounding Atlantic Assemian NE, Kouamé NG, Tohé B, Gourène G, Rödel Coast regions. PLoS ONE 9(10): e108213. MO. 2006. The anurans of the Banco National Park, Frost DR. 2015. Amphibian species of the World: An On- Côte d’Ivoire, a threatened West African rainforest. line Reference. Version 6.0. Available: http://research. Salamandra 42: 41‒51. amnh.org/vz/herpetology/amphibia (Accessed: 29 Assiri AA, Yoro GR, Deheuvels O, Kébé BI, Keli May 2015). American Museum of Natural History, ZJ, Adiko A, Assa A. 2009. Les caractéristiques New York, New York, USA. agronomiques des vergers de cacaoyer (Theobroma Grafe TU, Steffen JO, Stoll C. 2000. Vocal repertoire and cacao L.) en Côte d’Ivoire. Journal of Animal and effect of advertisement call intensity on calling behav- Plant Sciences 2: 55–66. iour in the West African tree frog, Leptopelis viridis. Bible M. 2013. The impacts of Côte d’Ivoire’s urbaniza- Amphibia-Reptilia 21: 13–23. tion on its economy and populace. Global Majority Greenbaum E, Carr JL. 2005. The herpetofauna of Upper E-Journal 4: 94–105. Niger National Park, Guinea, West Africa. Scientific Channing A, Howell KM 2006. Amphibians of East Afri- Papers of the Natural History Museum University of ca. Cornell University Press, Ithaca, New York, USA. Kansas 37: 1–21. 432 p. Hamer AJ, McDonnell MJ. 2008. Amphibian ecology Colwell RK. 2006. EstimateS, statistical estimation of and conservation in the urbanising world: A review.

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N’Goran Germain Kouamé is an Ivorian herpetologist and biologist. He is lecturer at the Jean Lorougnon University, Daloa, Ivory Coast and a member of the IUCN SSC Amphibian Specialist Group (ASG). He holds a Diploma and a Ph.D. in natural sciences from the University of Abobo-Adjamé, Abidjan, Ivory Coast, where he used leaf-litter frogs (Phrynobatrachus spp.) as models to determine the conservation status of the Banco National Park, one of the rare remaining primary forests situated in the midst of a West African mega-city. His current research interests focus on the taxonomy, ecology, distribution, and conservation of rare, threatened, and new amphibian species in Ivory Coast.

Caleb Ofori-Boateng is a research scientist at the Forestry Research Institute of Ghana. He holds a Ph.D. in wildlife management and a B.S. in Natural Resources Management from the Kwame Nkrumah University of Science and Technology in Ghana. His research focuses on ecology, population genetics and conservation of West African amphibians. Caleb is also the founder and Director of Herp Conservation Ghana (Herp-Ghana), a non-profit organization dedicated to amphibian and reptile conservation in West Africa.

Gilbert Baase Adum is a research scientist based at Ghana’s premiere science university Kwame Nkrumah Uni- versity of Science and Technology in Kumasi. He is also the co-founder and Executive Director of SAVE THE FROGS! Ghana, West Africa’s first non-profit organization dedicated exclusively to amphibian conservation. Through his work with SAVE THE FROGS! Ghana, Gilbert aims to help prevent the extinction of endangered frogs, while spreading the message of frog conservation and environmental protection across the entire African continent. He is currently working on a collaborative project at the Museum für Naturkunde, Berlin, with the aim of establishing knowledge about impacts of climate change on Ghanaian amphibians.

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Germain Gourène is the professor and founder of the “Laboratoire d’Environnement et de Biologie Aquatique” at the Nangui Abrogoua University (ex-University of Abobo-Adjamé, Abidjan, Ivory Coast). His research fo- cuses on the systematics and taxonomy of fishes with emphasis on Africa; areas covered also include ecology and aquaculture. In addition to his research interest on fishes Germain is interested in the conservation of aquatic invertebrates and amphibians in the Banco National Park. Germain has served as Vice-President and President of the University of Abobo-Adjamé for 10 years. He is embarking on a political career and has been elected as deputy of the locality of Kounahiri since 2012.

Mark-Oliver Rödel is the Curator of Herpetology and head of the department of “Diversity Dynamics” at the Museum für Naturkunde, Berlin, and teaches biodiversity at the Humboldt University, Berlin. Since his teenage age he has dedicated his life to the study of amphibians and reptiles, mostly to those from Africa. Mark-Oliver is the Chairman for West and Central Africa within the IUCN SSC Amphibian Specialist Group (ASG). With his team he investigates the taxonomy, systematics, and biogeography of amphibians and reptiles, but in particular uses amphibians as model organisms in order to understand the effect of environmental change on species and ecosystems.

Appendix 1. Geographic position and short description of study sites in the Daloa study area. Site Latitude (N) Longitude (W) Elevation (m a.s.l.) Habitat description Highway; grassy habitats; heavy traffic; dense human Bal N06°53’64.5” W006°25’65.3” 259 population Large stream bordered by coconut trees and grass; un- Eve N06°53’01.8” W006°26’05.9” 261 paved roads; concentration of buildings; swampy area used for vegetable cultivation; dense human population Semi-deciduous forest patch; swampy area dominated by grassy vegetation; buildings; highway, heavy traffic; Gbo 1 N06°54’15.8” W006°27’15.3” 265 streetlight; swampy area partly used for vegetable cultiva- tion; dense human population Buildings; shrubby vegetation; unpaved roads; highway; Gbo 2 N06°54’03.4” W006°27’09.6” 275 dense human population Subsistence farming; rice field in a swampy area; high Sap 1 N06°87’20.8” W006°37’83.8” 239 grassy vegetation Forests islands; cocoa plantation at edge of a rice field; Sap 2 N06°87’22.8” W006°37’93.7” 260 high grassy vegetation Stream crossing cocoa plantation; palm tree at edge of the Sap3 N06°87’37.1” W006°38’09.7” 276 water body; plantain and coffee plantations Swampy area; high grassy vegetation; rice field; humid Sap 4 N06°86’83.8” W006°38’99.3” 244 savanna; tracks Sap 5 N06°86’83.8” W006°37’61.5” 229 Rice field in a swampy area; coconut trees at edge Semi deciduous forest patch close to the Jean Lorougnon Guédé University; garden of the Theological and Pastoral Institute, dominated by bamboo, grasses and stands of dif- Taz 1 N06°90’42.8” W006°43’97.4” 274 ferent ornamental plants; two large wide ponds surrounded by vegetation (Asteraceae); streetlight; unpaved roads, concentration of buildings; dense human population Swampy area; buildings; rice field; vegetable cultivation; Taz 2 N06°90’33.7” W006°43’78.9” 268 many constructions of houses underway Swamps within a semi deciduous forest; stream; ponds; Zai 1 N06°94’97.8” W006°67’35.7” 223 grassy vegetation; clearing; rice field; cocoa plantation Very large rice field; high grasses; edge of coffee and Zai 2 N06°94’33.6” W006°68’89.0” 222 cocoa plantations; forest patch Rice field crossed by a stream: coffee plantation; tracks; Zai 3 N06°94’65.9” W006°66’58.7” 209 forest patch

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Appendix 2. Amphibian survey periods in the urban and non-urban areas of Daloa (compare Appendix 1). Surveyed periods Site 10–21 Aug. 27 Aug.–1 27 Sep.–1 16–25 Aug. 18–27 Oct. 18–27 Jun. 16–24 Aug. 2011 Sep. 2011 Nov. 2012 2013 2013 2014 2014 Urban area 6 days 2 days 1 day 6 days 2 days 6 days 3 days Bal 6 2 1 6 2 6 3 Eve 6 2 1 6 2 6 3 Gbo 6 2 1 6 2 6 3 Taz 6 2 1 6 2 6 3 Non-urban area 6 days 4 days 4 days 4 days 8 days 4 days 6 days Sap 3 2 2 2 4 2 3 Zai 3 2 2 2 4 2 3 Total days 12 6 5 10 10 10 9

Appendix 3. List of amphibian voucher specimens from the district of Daloa and surroundings. Given are field and collection num- bers (NG), collection site (compare Appendix 1) and collection date.

Arthroleptidae: Arthroleptis spp.: NG001 (Taz, 08 Nov. 2011); NG002 (Sap, 16 Oct. 2013); NG003 (Zai, 23 Oct. 2013); Leptopelis spiritusnoctis: NG004 (Zai, 23 Oct. 2013); L. viridis: NG005 (Taz, 24 Oct. 2011); NG006 (Taz, 30 Oct. 2013); Bufonidae: Ami- etophrynus maculatus: NG007 (Zai, 25 Oct. 2013); A. regularis: NG008 (Taz, 19 Oct. 2011); Dicroglossidae: Hoplobatrachus oc- cipitalis: NG009 (Taz, 25 Oct. 2011); Hemisotidae: Hemisus marmoratus: NG010 (Taz, 08 Sep. 2010); NG011 (Sap, 25 Oct. 2013); Hyperoliidae: Afrixalus dorsalis: NG012 (Taz, 18 Aug. 2011); Hyperolius concolor concolor: NG013 (Taz, 18 Aug. 2011); NG014 (Sap, 18 Oct. 2013); H. fusciventris fusciventris: NG015 (Sap, 19 Oct. 2013); H. guttulatus: NG016 (Zai, 23 Oct. 2013); NG017 (Zai, 25 Oct. 2013); H. nitidilus: NG018 (Taz, 25 Aug. 2011); NG019 (Sap, 16 Oct. 2013); H. picturatus: NG020 (Taz, 01 Sep. 2011); NG021 (Sap, 18 Oct. 2013); NG022 (Taz, 23 Oct. 2013); H. sp.: NG023 (Taz, 15 Sep. 2013); Kassina schioetzi: NG024–027 (Taz, 08 Sep. 2013); K. senegalensis: NG028 (Taz, 17 Aug. 2011); NG029 (Taz, 30 Aug. 2013); Microhylidae: Phrynomantis microps: NG030 (Taz, 08 Sep. 2013); Phrynobatrachidae: Phrynobatrachus calcaratus: NG031–036 (Sap, 19–20 Oct. 2013); P. francisci: NG037–038 (Taz, 01 Sep. 2011); NG039 (Taz, 17 Aug. 2013); P. gutturosus: NG040 (Taz, 19 Aug. 2013); NG041 (Sap, 20 Oct. 2013); P. latifrons: NG042 (Taz, 20 Aug. 2011); NG043 (Taz, 25 Aug. 2011); NG044 (Sap, 19 Oct. 2010); NG045 (Zai, 23 Oct. 2013); Ptychadenidae: Ptychadena bibroni: NG046 (Taz, 09 Sep. 2012); NG047 (Zai, 24 Oct. 2013); P. tellinii: NG048 (Taz, 01 Sep. 2013); P. mascareniensis: NG049 (Taz, 16 Sep. 2013); NG050 (Zai, 24 Oct. 2013); P. oxyrhynchus: NG051–052 (Taz, 06 Sep. 2012); NG053 (Zai, 24 Oct. 2013); P. pumilio: NG054 (Taz, 16 Sep. 2013); NG055 (Sap, 19 Oct. 2013); P. tournieri: NG056 (Taz, 10 Sep. 2012); Ranidae: Amnirana albolabris: NG057 (Sap, 19 Oct. 2013); A. galamensis: NG058 (Taz, 24 Aug. 2014).

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